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Factors Modulating Post-Activation Potentiation and its Effect on

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 Sports Med 2009; 39 (2): 147-166
REVIEW ARTICLE 0112-1642/09/0002-0147/$49.95/0

ª 2009 Adis Data Information BV. All rights reserved.

Factors Modulating Post-Activation

Potentiation and its Effect on Performance
of Subsequent Explosive Activities
Neale Anthony Tillin1,2 and David Bishop1,3
1 School of Human Movement and Exercise Science, the University of Western Australia, Crawley,
Western Australia, Australia
2 School of Sport and Exercise Science, Loughborough University, Loughborough, Leicestershire, UK
3 Facoltà di Scienze Motorie, Università degli Studi di Verona, Verona, Italy

Contents
Abstract. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
1. Post-Activation Potentiation (PAP). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
2. Mechanisms of PAP . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
2.1 Phosphorylation of Regulatory Light Chains. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
2.2 Increased Recruitment of Higher Order Motor Units . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
2.3 Changes in Pennation Angle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
3. PAP and Mechanical Power . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
4. Acute Effects of PAP on Subsequent Activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
4.1 PAP versus Fatigue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 156
4.2 Conditioning Contraction Volume . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
4.3 Conditioning Contraction Type . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
4.4. Subject Characteristics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
4.4.1 Muscular Strength . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
4.4.2 Fibre-Type Distribution. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
4.4.3 Training Level . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
4.4.4 Power-Strength Ratio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
4.5 Type of Subsequent Activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162
5. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163

Abstract Post-activation potentiation (PAP) is induced by a voluntary conditioning

contraction (CC), performed typically at a maximal or near-maximal in-
tensity, and has consistently been shown to increase both peak force and rate
of force development during subsequent twitch contractions. The proposed
mechanisms underlying PAP are associated with phosphorylation of myosin
regulatory light chains, increased recruitment of higher order motor units,
and a possible change in pennation angle. If PAP could be induced by a CC in
humans, and utilized during a subsequent explosive activity (e.g. jump or
sprint), it could potentially enhance mechanical power and thus performance
and/or the training stimulus of that activity. However, the CC might also
induce fatigue, and it is the balance between PAP and fatigue that will
148 Tillin & Bishop

determine the net effect on performance of a subsequent explosive activity.

The PAP-fatigue relationship is affected by several variables including CC
volume and intensity, recovery period following the CC, type of CC, type of
subsequent activity, and subject characteristics. These variables have not
been standardized across past research, and as a result, evidence of the effects
of CC on performance of subsequent explosive activities is equivocal. In or-
der to better inform and direct future research on this topic, this article will
highlight and discuss the key variables that may be responsible for the con-
trasting results observed in the current literature. Future research should aim
to better understand the effect of different conditions on the interaction be-
tween PAP and fatigue, with an aim of establishing the specific application
(if any) of PAP to sport.

1. Post-Activation Potentiation (PAP) of factors that influence acute performance fol-

Post-activation potentiation (PAP) or post-
tetanic potentiation (PTP) refers to the phenomena
by which muscular performance characteristics
are acutely enhanced as a result of their contrac-
tile history.[1,2] The difference between PAP and
PTP is defined by the nature of the conditioning
contraction. PTP is induced by an involuntary
tetanic contraction, and PAP is induced by a
voluntary contraction[3,4] performed typically
at a maximal or near-maximal intensity. For
simplicity, this article refers to all potentiation
responses as PAP, and refers to the activity res-
ponsible for inducing PAP as a conditioning
contraction (CC).
The presence of PAP in skeletal muscle has
been recorded by many studies in both mammals
and humans,[5-17] prompting a discussion
amongst recent review articles over the mechan-
isms of PAP[1,3] and its application to sports
performance.[1-3,18] If effectively utilized, PAP
could be implemented into a power-training
routine to enhance the training stimulus of a
plyometric exercise.[2,18] Inducing PAP prior to
competition might also prove better than con-
ventional warm-up techniques at enhancing per-
formance of explosive sports activities such as
jumping, throwing and sprinting.[10] Because of
inconsistencies within the literature, research re-
mains inconclusive on the possible benefits of
PAP to explosive sports performance and/or
training. The inconsistencies of past research
are most likely due to the complex interaction
lowing a CC.[1-3,18] This review discusses these
confounding factors in greater detail, with the
purpose of helping to inform and direct future
research efforts towards establishing the appli-
cation (if any) of PAP to performance/training of
explosive sports activities.
2. Mechanisms of PAP
It has been proposed that two principal me-
chanisms are responsible for PAP. One is the
phosphorylation of myosin regulatory light
chains (RLC),[1,3,4,11,12,19,20] and the other is an
increase in the recruitment of higher order motor
units.[1,10,20] There is also evidence to suggest that
changes in pennation angle may contribute to
PAP, and this possible mechanism is briefly in-
troduced in this article.
2.1 Phosphorylation of Regulatory Light
Chains
A myosin molecule is a hexamer composed of
two heavy chains (figure 1).[21] The amino-
termini of each heavy chain, classified as the
myosin head, contain two RLCs,[9,21] and each
RLC has a specific binding site for incorporation
of a phosphate molecule. RLC phosphorylation
is catalyzed by the enzyme myosin light chain ki-
nase, which is activated when Ca2+ molecules,
released from the sarcoplasmic reticulum dur-
ing muscular contraction, bind to the calcium
regulatory protein calmodulin.[1,5,13,21] RLC

ª 2009 Adis Data Information BV. All rights reserved. Sports Med 2009; 39 (2)
Post-Activation Potentiation, Theory and Application
Actin binding site
RLC-2
Myosin heavy chains
ATP binding site
Fig. 1. One myosin molecule. Each myosin molecule is composed
of two myosin heavy chains. Regulatory light chain (RLC)-2 re-
presents a pair of RLCs positioned at the neck of a myosin head.
Each RLC can incorporate a phosphate molecule, altering the
structure of the myosin head. At each myosin head there is an actin
and adenosine triphosphate (ATP) binding site.
phosphorylation is thought to potentiate sub-
sequent contractions by altering the structure
of the myosin head and moving it away from
its thick filament backbone.[1,21] It has also
been shown that RLC phosphorylation renders
the actin-myosin interaction more sensitive to
myoplasmic Ca2+.[13] Consequently, RLC phos-
phorylation has its greatest effect at relatively
low concentrations of Ca2+, as is the case
during twitch or low-frequency tetanic
contractions.[1,3,4,22,23]
An acute increase in RLC phosphorylation,
and a parallel potentiation of twitch tension fol-
lowing tetanic stimulation of specific efferent
neural fibres, has been reported by many studies
in skinned animal models[5,7,9,13] (figure 2). Re-
latively few studies have attempted to measure a
similar response in human skeletal muscle. Stuart
et al.[8] recorded a significantly elevated phos-
phate content of RLC in the vastus lateralis
muscle (p < 0.01), and a significant potentiation of
twitch tension of the knee extensors, following
one 10-second isometric maximal voluntary con-
traction (MVC; p < 0.05). There was also a posi-
tive but non-significant correlation between the
extent of twitch potentiation and the amount of
phosphate incorporated into individual RLC
units, and between potentiation and percentage
of type II muscle fibres (p > 0.05).
Smith and Fry[24] also sampled muscle biopsies
at the vastus lateralis, and analysed dynamic
leg extension performance before and 7 minutes
after a 10-second isometric MVC. The authors
ª 2009 Adis Data Information BV. All rights reserved.
149
reported no significant change in RLC phos-
phorylation or leg extension performance for the
entire sample (p > 0.05). The subjects were then
split into those who responded to the MVC with a
significant increase, and those who responded
with a significant decrease in RLC phosphoryla-
tion (p < 0.05), but no significant differences in leg
extension performance were found between the
groups (p > 0.05). Methodological factors and
differences in fibre-type distribution between an-
imals and humans may explain why an observed
increase in RLC phosphorylation following a
CC is not as consistent in humans as animals.
Nevertheless, the significance of RLC phosphory-
lation in human skeletal muscle remains unclear,
and Stuart et al.[8] suggest that other factors may
provide the major contribution to PAP.
2.2 Increased Recruitment of Higher Order
Motor Units
Research on animals has shown that an in-
duced tetanic isometric contraction (caused by
stimulating specific afferent neural fibres, which
in turn activate adjacent a-motoneurons via an
afferent neural volley; figure 3) elevates the
transmittance of excitation potentials across
synaptic junctions at the spinal cord. This ac-
commodating state can last for several minutes
following the tetanic contraction,[10] and as a
Phosphate content
0.6 2.0
mol phosphate/mol RLC
potentiation (post/pre)
Twitch potentiation
Twitch peak torque
1.8
0.4
1.6
1.4
0.2
1.2
0 1.0
0 10 20 70 130 190 250
Time (sec)
Tetanic
contraction
Fig. 2. The time-course of regulatory light chain (RLC) phosphor-
ylation and twitch peak torque potentiation, following a 10-second
pre-conditioning tetanus. Potentiation is represented as a ratio of the
post-maximal voluntary contraction (MVC) peak torque value to the
pre-MVC peak torque value (post/pre). These results indicate
a possible relationship between RLC phosphorylation and twitch
tension potentiation (reproduced from Moore and Stull,[7] with
permission).
Sports Med 2009; 39 (2)
150
Alpha motoneuron
synapse
Spinal cord
Alpha motoneuron Alpha motoneuron
to agonist
to synergist
Afferent neural Alpha motoneuron
fibre (la)
to antagonist
Muscle spindle Antagonist
muscle
Agonist muscle
Synergist
muscle
Fig. 3. The neural volleys of a Ia afferent fibre. An action potential
generated at the Ia afferent neural fibre travels to the spinal cord,
where it is transferred to the adjacent a-motoneuron of the agonist
muscle. The action potential then travels directly to the agonist
muscle, initiating the processes of muscular contraction.
result there is an increase in post-synaptic po-
tentials, for the same pre-synaptic potential dur-
ing subsequent activity.[25,26]
Luscher et al.[26] proposed a possible mechan-
ism underlying the elevated transmittance of ac-
tion potentials across synaptic junctions at the
spinal cord. For each parent neural fibre (i.e. Ia
fibre) numerous synapses project onto each
a-motoneuron. Activation of an a-motoneuron
works in an all-or-none fashion, whereby pre-
synaptic transmitter release must coincide with
the post-synaptic receptor sensibility. Transmit-
ter failure at various synaptic junctions is a
common occurrence during normal reflex or vo-
luntary responses, due to an autonomously pro-
tected activation reserve.[26,27] An induced tetanic
contraction is suggested to decrease the trans-
mitter failure during subsequent activity, via one
or a combination of several possible responses.
These include an increase in the quantity of neuro-
transmitter released, an increase in the efficacy
of the neurotransmitter, or a reduction in axonal
branch-point failure along the afferent neural
fibres.[28]
ª 2009 Adis Data Information BV. All rights reserved.
Tillin & Bishop
Hirst et al.[27] provided evidence to support a
decreased monosynaptic transmitter failure dur-
ing subsequent activity. They stimulated cat af-
ferent neural fibres, and observed a 54% increase
in excitatory post-synaptic potentials (EPSPs)
for the same pre-synaptic stimulus, following a
20-second tetanic isometric contraction. Larger
EPSPs represent greater depolarization of the
a-motoneuron membrane, which would increase
the likelihood of that a-motoneuron reaching the
threshold required to initiate an action potential,
and subsequently contract the muscle fibres of
that motor unit.
Luscher et al.[26] also measured EPSPs at cat
a-motoneurons, in response to electrical stimu-
lation. They found a significant positive correla-
tion between motoneuron input resistances and
EPSP amplitude, for a standard stimulus (r = 0.77;
p < 0.01; figure 4a), where input resistance was
associated with the size of the a-motoneuron
(with a smaller input resistance representing a
larger motoneuron). This suggests that mono-
synaptic transmitter failure is greater at larger
motoneurons (those responsible for activation
of higher order or fast-twitch motor units). Con-
versely, when a twitch was stimulated following a
10-second tetanic contraction, Luscher et al.[26]
found a significant negative correlation between
EPSP potentiation and motoneuron input
resistance (r = -0.92; p < 0.001; figure 4b). This
demonstrates that a tetanic contraction decreased
the transmitter failure occurring primarily
at larger motoneurons, which resulted in a
considerable PAP effect at these motoneurons.
If a CC could induce an increase in higher order
motoneuron recruitment in humans, this effect
might theoretically increase fast-twitch fibre
contribution to muscular contraction, and there-
fore enhance performance of a subsequent ex-
plosive activity.[10]
Previous studies have measured the H-wave in
humans to investigate the effects of a CC on
motoneuron recruitment.[10,29] The H-wave
(H-reflex) is recorded at the muscle fibres using
electromyography, and is the result of an afferent
neural volley in response to single-pulse sub-
maximal stimulation of the relevant nerve bundle
(see figure 5 for more detail). An increase in
Sports Med 2009; 39 (2)
Post-Activation Potentiation, Theory and Application
H-wave following a CC may therefore represent
a decrease in transmitter failure at synaptic
junctions, and a subsequent increase in higher
order motoneuron recruitment. Gullich and
Schmidtbleicher[10] stimulated the tibial nerve
and measured changes in H-wave amplitude at
the gastrocnemius before and after five 5-second
isometric MVCs of the plantarflexors. They re-
ported a depression in H-wave amplitude 1 minute
a
12
% Increase in EPSP amplitude
6 an action potential travelling directly down the efferent neural fibres
0
0 1 2 3
Larger motoneurons Smaller motoneurons
Input resistance (MΩ)
b was not normalized to maximal M-wave (M-wave
140
% Increase in EPSP amplitude
70
0 and 5–11 minutes post a 10-second isometric
0 1 2 3
Larger motoneurons Smaller motoneurons
Input resistance (MΩ)
Fig. 4. (a) The relationship between input resistances of cat moto-
neurons, and amplitude of their excitatory post-synaptic potentials
(EPSP) in response to twitch stimulation of the adjacent afferent
neural fibres. (b) The relationship between input resistances of cat
motoneurons, and the percentage increase (potentiation) in EPSP
amplitude, in response to a twitch stimulation of the adjacent afferent
neural fibres, following a 10-second tetanus. Although EPSP ampli-
tude is greatest at smaller motoneurons (those with greater input
resistances), representing greater transmitter failure at larger moto-
neurons (a), potentiation is greatest at larger motoneurons (those
with smaller input resistances), demonstrating a decreased trans-
mitter failure at these motoneurons (b).[22]
ª 2009 Adis Data Information BV. All rights reserved.
151
1st response to the electrical
stimulation (M-wave)
Muscle
Spinal cord Afferent neural fibres
Electrical
stimulation
Efferent neural fibres
2nd response to the electrical
stimulation (H-wave)
Fig. 5. Elicitation of an M- and H-wave. Stimulation of a nerve with
a single submaximal electrical impulse evokes two electrical re-
sponses at the muscle. The first response (M-wave) is the result of
(a-motoneurons). The second response (H-wave) is the result of an
action potential travelling along the afferent neural fibres to the spinal
cord, where it is transmitted to adjacent efferent neural fibres, and
down to the muscle.
4 5 after the MVCs (-24%; p < 0.05), but a potentia-
tion of H-wave amplitude 5–13 minutes after the
MVCs (+20%; p < 0.01). The H-wave, however,
is the electrical counterpart of the activation of all
motor units in the pool[30]). Therefore, other
factors not relating to central activation, such as
increased activity of the Na+-K+ pump at the
muscle fibres,[12,14,28] may be responsible for the
results that Gullich and Schmidtbleicher[10] ob-
served. Nevertheless, other studies have reported
a potentiation in normalized H-wave amplitude
3–10 minutes post eight sets of dynamic MVCs,[29]
4 5 MVC.[31] Collectively, these results suggest that
PAP increases H-wave amplitude in humans
(albeit after sufficient recovery), and this may be
the result of increased higher order motoneuron
recruitment at the spinal cord. Whether or not a
CC can enhance motoneuron recruitment and
performance during a subsequent voluntary con-
traction is yet to be determined.
The effect of isometric MVCs on subsequent
voluntary motoneuron recruitment has been as-
sessed using the interpolated twitch technique
(ITT). The ITT can facilitate measurement of
Sports Med 2009; 39 (2)
152
motoneuron activation[32] by comparing maximal
twitch amplitude at rest with that evoked when
superimposed upon an MVC (for more detail of
the ITT please refer to Folland and Williams[32]
and Shield and Zhou[33]). Using the ITT, Behm
et al.[34] reported a decrease in voluntary muscle
activation following 10-second MVCs (p < 0.05).
These results are in contrast to the proposed
mechanism of PAP, but may demonstrate the
dominance of central fatigue observed through-
out this study (see section 4.2). Nevertheless, fu-
ture research should consider using the ITT to
investigate the mechanisms of PAP and their
contribution to subsequent performance.
2.3 Changes in Pennation Angle
The pennation angle of a muscle (the angle
formed by the fascicles and the inner apo-
neurosis) reflects the orientation of muscle fibres
in relation to connective tissue/tendon.[35] The
pennation angle will therefore affect force trans-
mission to the tendons and bones.[35,36] The sum
of the forces of all individual fibres being applied
to the relevant tendon during muscular contrac-
tion is reduced by a factor of cosy (where y =
pennation angle).[36] Consequently, smaller pen-
nation angles have a mechanical advantage with
respect to force transmission to the tendon.[35,36]
Using ultrasonography, Mahlfeld et al.[37] mea-
sured resting pennation angle of the vastus la-
teralis before and after three 3-second isometric
MVCs. Pennation angle immediately after the
MVCs (15.7) had not changed from pre-MVC
values (16.2); however, 3–6 minutes after the
MVCs, the pennation angle had significantly de-
creased (14.4; p < 0.05). This change would only
be equivalent to a 0.9% increase in force trans-
mission to the tendons, but it is possible that this
effect may contribute to PAP. Conditioning
contractions, however, are also likely to increase
connective tissue/tendon compliance,[38] and this
may counter any increase in force transmission
caused by a decrease in pennation angle. Never-
theless, the possibility that changes in muscle
architecture contribute to PAP warrants further
investigation.
ª 2009 Adis Data Information BV. All rights reserved.
Tillin & Bishop
3. PAP and Mechanical Power
Performance of explosive sports activities is
largely determined by mechanical power.[10,39-43]
Mechanical power can be defined as the rate
at which force (F) is developed over a range
of motion (d), in a specific period of time (t)
[P = F · d/t], or as force multiplied by velocity (v)
[P = F · v].[39,40,43] Accordingly, increasing the
level of force at a given velocity will increase
mechanical power, and this has been demon-
strated in skinned rat/mouse models.[16,17,22]
Similarly, decreasing the time over which a speci-
fic force is applied, without altering the distance
over which that force is applied, will increase
velocity, and consequently mechanical power.
PAP could, therefore, increase force and/or
velocity of the muscle contraction, which would
enhance mechanical power and the associated
sport performance.
To date, there is little evidence that PAP can
increase maximal force. This is consistent with
the observation that increased sensitivity of the
myosin-actin interaction to Ca2+ has little or no
effect in conditions of Ca2+ saturation, such as
those caused by higher stimulation frequencies
(>20 Hz for tetanic, or 200 Hz for voluntary
contractions).[9,22] Stuart et al.[8] also found that a
10-second isometric MVC of the knee extensors
was unable to increase maximum unloaded velo-
city of subsequent dynamic contractions. Al-
though PAP appears to have little effect at the
extremes of the force-velocity curve (figure 6), it
has been shown to increase rate of force devel-
opment (RFD) of tetanic contractions elicited at
any frequency.[9] An increase in RFD causes a
less concave force-velocity curve (figure 6), re-
sulting in a greater velocity for a specific force, or
vice versa.[3,44] Therefore, PAP may enhance the
performance of activities that require sub-
maximal force and velocity production.[3,11]
Typically, athletes participating in explosive
sports activities will not produce maximal force
because the mass they are attempting to move is
often relatively small (e.g. body mass), but they
must still overcome that mass so will not achieve
maximal unloaded velocity either.[40] Conse-
quently, PAP could benefit the performance
Sports Med 2009; 39 (2)
Post-Activation Potentiation, Theory and Application
100
Percentage of maximum unloaded velocity
Increased RFD
0 at the femoral nerve prior to, 5 seconds after, and
0 100
Percentage of maximum force
Fig. 6. The relationship between force and velocity. The dotted line
represents a less concave force-velocity curve due to an increase
in rate of force development (RFD) [reproduced from Sale,[3] with
permission].
of explosive sports activities by increasing RFD
and thus mechanical power.[3,11]
There is consensus over the existence of PAP,
but if it is to be effectively utilized in performance
and/or training, research must first confirm that
PAP can be induced by an isometric or dynamic
voluntary contraction, and then show that its
benefits can be realized during a subsequent ex-
plosive sports activity. Unfortunately, measure-
ment of both PAP and its effect on performance
of a subsequent explosive sports activity in
humans is inconsistent. Furthermore, little is
known about the degree to which the proposed
mechanisms underlying PAP may play a role in
inducing an elevated neuromuscular response.
4. Acute Effects of PAP on Subsequent
Activity
The performance of explosive sports activities
relies predominantly on the activation of large
muscle groups (e.g. ankle, knee, hip and/or arm
and ab/adductors). Therefore, studies assessing
the effect of PAP on smaller muscle groups have
been excluded from the following sections. Fur-
thermore, it has been shown[45,46] and is widely
accepted that contractions of maximal or near
maximal intensity (>80% of dynamic or iso-
metric MVC) optimize PAP.[4] Therefore, studies
ª 2009 Adis Data Information BV. All rights reserved.
153
assessing the effects of low-intensity contractions
on subsequent performance have also been
excluded from the following sections. Table I
summarizes the studies that have investigated
the effects of a voluntary CC on subsequent
voluntary activity in humans.
In agreement with the results produced by
studies conducted on skinned mammalian
models, research has consistently reported an
enhanced twitch response following a CC in
humans. Hamada et al.[12] elicited a twitch reflex
then every 30 seconds for 300 seconds after a
10-second isometric MVC of the knee extensors.
Twitch Pt (peak torque) was significantly in-
creased 5 seconds after the isometric MVC
(+71%; p < 0.01); however, by 30 and 60 seconds
after the isometric MVC, twitch Pt potentiation
had decreased to +44% and +31%, respectively
(p < 0.01). Potentiation continued to decrease at a
more gradual rate for the remainder of the re-
covery period, but was still +12% 300 seconds
after the isometric MVC (p < 0.01). Similar find-
ings have been reported in other studies,[6,11,59]
demonstrating that peak PAP is achieved im-
mediately after a CC, but instantly begins to de-
crease. The decrease in PAP is rapid for the first
minute, but then becomes more gradual resem-
bling an exponential function (figure 7).
Although an isometric MVC has been found
to consistently enhance subsequent twitch ten-
sion, evidence to show that PAP can be effectively
utilized to enhance the performance of sub-
sequent voluntary contractions is not as convin-
cing. Gossen and Sale[11] assessed movement
mechanics of both twitch and submaximal vo-
luntary contractions following a 10-second iso-
metric MVC. While the MVC enhanced twitch Pt
(p < 0.01), knee extension peak velocity follow-
ing the MVC was significantly lower than knee
extension peak velocity executed in a control
condition (326.7 vs 341.6/sec; p < 0.03). These
results suggest that although the 10-second MVC
induced PAP, it also induced fatigue, and that
the latter was more dominant during the volun-
tary contractions. It has been proposed, there-
fore, that it is the balance between PAP and
fatigue that determines whether the subsequent
Sports Med 2009; 39 (2)
ª 2009 Adis Data Information BV. All rights reserved.

154
Table I. A summary of studies that have investigated the effects of a pre-conditioning contraction on a subsequent activity
Study Subjects Pre-conditioning contraction Volume Rest interval Performance test Performance changes
(condition)
Batista et al.[47] 10 UT M Isovelocity MVC, knee 10 (30 sec RI) 4 min Isovelocity knee 6% › Pt* at each rest
extension 6 min extensions at all rest interval
8 min intervals
10 min

Behm et al.[34] 9 UT M Isometric MVC, knee 1 · 10 sec 1, 5, 10, 15 min Isometric MVC knee 2
extension 2 · 10 sec for all volumes extensions at all rest 2
(1 min RI) intervals 10-min post: 8.9% fl Pf*
3 · 10 sec 15-min post: 7.5% fl Pf *
(1 min RI)

Chatzopoulos et al.[48] 15 UT M Back-squat 10 · 1 rep 90% 3 min 30-m sprint 2

1 RM (3 min RI) 5 min 30-m sprint 3% fl 0–10-m sprint time*,
2% fl 0–30-m sprint time*

Chiu et al.[20] 24; 7 RT, 17 UT Back-squat 90% 1 RM · 5 (2 min RI) 5 min CMJ: 30% 1 RM RT: 1–3% › , UT: 1–4% fl .
(12 M, 12 F) 6 min 50% 1 RM 70% 1 RM RT > UT*
7 min SJ: RT: 1–3% › , UT: 1–4% fl .
5 min 30% 1 RM RT > UT*
6 min 50% 1 RM RT: 1–3% › , UT: 1–4% fl .
7 min 70% 1 RM RT = UT
RT: 1–3% › , UT: 1–4% fl .
RT > UT*
RT: 1–3% › , UT: 1–4% fl .
RT = UT
RT: 1–3% › , UT: 1–4% fl .
RT = UT

Ebben et al.[49] 10 RT M Dynamic bench-press 3–5 RM 0–5 sec Medicine ball BPT 2 GRF

French et al. [50]

14 RT (10 M, Isometric MVC, knee 3 sec · 3 (3 min RI) 0–5 sec CMJ 2
4 F) extension 5 sec · 3 (3 min RI) DJ 5.0% › * (4.9% › GRF*)
5 sec C-sprint 2 6.1% › Pt * 2
Isovelocity KE CMJ DJ 2
5 sec C-sprint isovelocity 2
KE 3.0% fl Pt *

Gilbert et al.[51] 7 RT M Back-squat 100% 1 RM · 5 (5 min RI) 2 min Isometric MVC at all rest 5.8% fl RFD
10 min intervals 5.8% fl RFD
15 min 10.0% › RFD
Sports Med 2009; 39 (2)

20 min
13.0% › RFD*
30 min
2

Tillin & Bishop

Gossen and Sale[11] 10 UT (6 M, Isometric MVC, knee 10 sec 20 sec Dynamic KE 2
4 F) extension 40 sec Dynamic KE 2

Continued next page

 Table I. Contd
ª 2009 Adis Data Information BV. All rights reserved.

Post-Activation Potentiation, Theory and Application

Study Subjects Pre-conditioning contraction Volume Rest interval Performance test Performance changes
(condition)
Gourgoulis et al.[15] 20 M (11 RT, Back-squats 2 reps of: 20%, 40%, 0–5 sec CMJ 2.4% › RT + UT*
9 UT) 60%, 80%, and 90% 1RM RT: 4.0% ›
(5 min RI) UT: 0.4% ›

Gullich and Study 1: 34 RT Isometric MVC, leg press 3 · 5 sec (5 min RI) 3 min, then every CMJ and DJ 3.3% › CMJ*. › DJ*
Schmidtbleicher[10] (22 M, 12 F) Isometric MVC, plantarflexion 5 · 5 (1 min RI) 20 sec. 8 jumps Isometric MVC, 13% fl RFD 1 min post*.
Study 2: 8 RT measured plantarflexion RFD 3 min post. 19% ›
1 min, then every 2nd RFD 5–13 min post*
min for 13 min

Hanson et al.[52] 30 UT (24 M, Back-squats 4 reps of 80% 1 RM 5 min CMJ 2

6 F)

Jenson and Ebben[53] 21 RT (11 M, Back-squats 5 RM 10 sec CMJ 4–13% fl *

10 F) 1 min CMJ 2
2 min CMJ 2
3 min CMJ 2
4 min CMJ 2

Kilduff et al.[54] 23 RT M Dynamic back-squats 1 · 3RM 15 sec CMJ 2.9% fl Pp*

Dynamic bench-press 1 · 3 RM 4 min CMJ 2
8 min CMJ 6.8% › Pp*
12 min CMJ 8.0% › Pp *
16 min CMJ 2
20 min CMJ 2
15 sec Barbell BPT
4.7% fl Pp *
4 min Barbell BPT
2
8 min Barbell BPT
2.8% › Pp*
12 min Barbell BPT
16 min Barbell BPT 5.3% › Pp*
20 min Barbell BPT 0.8% › Pp*

Magnus et al.[55] 10 UT M Back-squats 90% 1 RM 3 min CMJ 2

Rahimi[45] 12 RT M Back-squats 2 · 4 reps of 80% 4 min 40-m sprint 3% fl 0–40 m sprint time*
1 RM (2 min RI)

Rixon et al.[56] 30 UT (15 M, Dynamic back-squats 3 RM 3 min CMJ 2.9% › JH *, 8.7% › Pp *

15 F) Isometric MVC back-squats 3 · 3 sec (2 min RI) 3 min CMJ 2 JH, 8.0% › Pp *

Robbins and 16 UT M Isometric MVC back-squats 3 · 7 sec (8 min 4 min CMJ after each set of 2
Sports Med 2009; 39 (2)

Docherty[57] between each set) isometric MVC

Young et al.[58] 10 UT M Back-squats 5 RM 4 min LCMJ 2.8% › *

BPT = bench press throw; CMJ = counter movement jump; C-sprint = cycle sprint; DJ = drop jump; F = females; GRF = ground reaction force; JH = jump height; KE = knee extensions;
LCMJ = loaded counter movement jump; M = males; MVC = maximum voluntary contractions; Pf = peak force; Pp = peak power; Pt = peak torque; RFD = rate of force development;
RI = rest interval; RM = repetition maximum; RT = resistance/athletically trained; SJ = squat jump; UT = un/recreationally trained; › indicates increase; fl indicates decrease;

155
2 indicates no differences; * p < 0.05.
156 Tillin & Bishop
Twitch peak torque potentiation (post/pre)

1.8 shows how the net affect on subsequent volun-

1.7 tary contractions might be very different to the
1.6
effect of a MVC on subsequent twitch contrac-
tions (represented in figure 7).
1.5
There is also evidence that a recovery period
1.4 may not be required to benefit from PAP, or that
1.3 even with a recovery period performance of
1.2 a subsequent voluntary activity may remain
unchanged/diminished. French et al.[50] did not
1.1
utilize a recovery period, but still observed a sig-
1.0 nificant increase in both drop jump (DJ) height
0 30 60 90 120 150 180 210 240 270 300 and isovelocity knee extension Pt (+5.0% and
Time immediately after a 10-sec isometric MVC (sec) +6.1%, respectively; p < 0.05), immediately after
Fig. 7. The time-course of twitch peak torque potentiation im- three sets of 3-second isometric MVC knee ex-
mediately after a 10-second isometric maximal voluntary contraction tensions. Likewise, Gourgoulis et al.,[15] reported
(MVC).[12] Potentiation is represented as a ratio of the post-MVC
peak torque value to the pre-MVC peak torque value (post/pre). a significant increase in CMJ height (+2.4%;
p < 0.05) immediately after two back-squats per-
contractile response is enhanced, diminished or formed with 90% of one repetition maximum
unchanged.[2] (1RM). Conversely, Chiu et al.[20] were unable to
detect a significant improvement in peak power
4.1 PAP versus Fatigue of three CMJs or three loaded squat jumps (SJ)
[p > 0.05], even though they were performed after
The balance between PAP and fatigue and its a recovery period of 5, 6 and 7 minutes, respec-
effect on subsequent explosive contractions has tively, following five sets of one back-squat, with
been observed by several studies. Immediately 90% 1RM. The three CMJs (5, 6 and 7 minutes
after a CC, Gullich and Schmidtbleicher[10] and post-activation), were executed with different
Gilbert et al.[51] reported a decrease or no change
in isometric RFD, but following a sufficient re-
covery (4.5–12.5 minutes[10] and 15 minutes[51]) Peak PAP
isometric RFD was significantly increased 2
(+10–24%; p < 0.05). The same pattern of no
change/decrease followed by an increase in
Potentiation (post/pre)

Performance
counter-movement jump (CMJ) peak power
(+7–8%; p < 0.05)[54] and 30-m sprint perfor-
mance (2–3%; p < 0.05)[48] 8–12 minutes and 1
5 minutes, respectively, following a CC have also
been reported. Collectively, these results suggest
Peak fatigue
that although twitch studies have reported max- Window
imal PAP immediately after a CC (described in 1 Window 2
0
section 4; see figure 7), fatigue is also present Condition volume Recovery time
early on. Furthermore, fatigue seems more
dominant in the early stages of recovery and, Fig. 8. A model of the hypothetical relationship between post-
activation potentiation (PAP) and fatigue following a pre-conditioning
consequently, performance of subsequent volun- contraction protocol (condition).[3] When the condition volume is low,
tary activity is diminished or unchanged. How- PAP is more dominant than fatigue, and a potentiation in subsequent
explosive performance (post/pre) can be realized immediately (win-
ever, fatigue subsides at a faster rate than PAP, dow 1). As the condition volume increases, fatigue becomes domi-
and potentiation of performance can be realized nant, negatively affecting subsequent performance. Following the
condition, fatigue dissipates at a faster rate than PAP, and a po-
at some point during the recovery period. Figure 8 tentiation of subsequent explosive performance can be realized at
illustrates the PAP-fatigue relationship and some point during the recovery period (window 2).

ª 2009 Adis Data Information BV. All rights reserved. Sports Med 2009; 39 (2)
Post-Activation Potentiation, Theory and Application 157

loads (30%, 50% and 70% of 1RM, respectively),
which may have affected peak power output, and
makes it difficult to compare differences in perfor-
mance over the time-course. However, these re-
sults were supported by those of Mangus et al.,[55]
who reported no change in CMJ height 3 minutes
after one back-squat with 90% 1RM. Finally,
Behm et al.[34] observed no change in isometric
peak force immediately after three 10-second MVCs;
however, after a 10- to 15-minute recovery period,
maximal force had decreased (7–9%; p < 0.05).
These contradictory findings suggest that the
PAP-fatigue relationship and its effects on sub-
sequent voluntary activity are multi-faceted.
In summary, it has been suggested that fol-
lowing a CC an optimal recovery time is required
to diminish fatigue and realize PAP; however,
evidence is inconsistent in support of this theory.
There are a number of possible explanations for
the contrasting results produced by the afore-
mentioned studies. The relationship between
PAP and fatigue, and the overall effect of con-
tractile history on subsequent performance, is
influenced by a combination of factors.[2] These
include: volume of the CC (e.g. sets, repetitions
and rest interval between numerous sets); in-
tensity of the CC (although there is consensus
that maximal-intensity contractions optimize
PAP), the type of CC performed (e.g. dynamic or
isometric); subject characteristics (e.g. muscular
strength, fibre-type distribution, training status
or power-strength ratio), and the type of activity
performed after the CC.[1,2] Figure 9 illustrates
the interaction of these complex factors and the
following sections discuss them in more detail.
4.2 Conditioning Contraction Volume
The effect of the CC volume on the interaction
between PAP and fatigue is highlighted by one
particular study. Hamada et al.[14] used a fati-
guing protocol of 16 5-second isometric MVC
knee extensions, with each MVC separated by
a 3-second rest interval. A twitch response was
stimulated at the femoral nerve pre-MVCs, bet-
ween each MVC, 1 minute after the MVCs, and
then every second minute after the MVCs, for
13 minutes. Twitch Pt gradually augmented over
the first three MVCs, peaking at a 127% increase
from baseline values (p < 0.05). This demon-
strates that PAP was more dominant than fati-
gue, after the first three MVCs when the MVC
volume was small. For the remainder of the fati-
gue protocol, however, twitch Pt progressively
decreased, and by the sixteenth MVC measured
32% below baseline-values (p < 0.05). This de-
monstrates that as the volume of MVCs con-
tinued to increase, so did the dominance of
fatigue. Following the fatigue protocol twitch Pt
gradually increased, and exceeded baseline values
after 30–120 seconds of recovery (+32%; p < 0.05).
This demonstrates that fatigue dissipated at a
faster rate than PAP and, consequently, there was
a potentiation in twitch Pt during the recovery

Condition Mechanisms of PAP:

intensity • RLC phosphorylation
• ↑ higher order
Subject characteristics:
motor-unit
Condition • Muscular strength Type of
recruitment Recovery
volume • Fibre type distribution explosive Performance
• ↓ pennation angle time
• Training level activity
• Strength-power ratio
Condition
type: Mechanisms of fatigue:
• Dynamic • Central
• Isometric • Peripheral

Fig. 9. The complex factors influencing performance of a voluntary explosive activity following a conditioning contraction (condition). Con-
dition intensity, volume and type will affect individuals differently, depending on their subject characteristics. Collectively, these factors will
influence the extent to which the mechanisms of post-activation potentiation (PAP) and fatigue are affected. The interaction between the
mechanisms of PAP and fatigue will determine whether subsequent performance is potentiated, and the recovery period required to realize
potentiation. Regardless of the previous interactions, however, the response of some explosive activities to the condition may be different to
the response of other explosive activities. RLC = regulatory light chain.

ª 2009 Adis Data Information BV. All rights reserved. Sports Med 2009; 39 (2)
158
period. An adaptation of these results is presented
in figure 10. These findings were supported in
another study.[6] They recorded twitch tension
in the dorsiflexors before and immediately
after five isometric dorsiflexion MVC protocols,
where each protocol differed in MVC dura-
tion (volume). Accordingly, each protocol in-
duced a different level of PAP, with a 10-second
isometric MVC eliciting the greatest potentia-
tion (twitch Pt: after a 1-second MVC = +43%;
after a 3-second MVC = +130%; after a 10-second
MVC = +142%; after a 30-second MVC = +65%;
after a 60-second MVC = +14%). Again, the
important question is whether or not a similar
effect will occur during performance of voluntary
explosive activities?
French et al.[50] assessed the effect of different
CC volumes on performance of subsequent vo-
luntary explosive activities. They measured a
significant increase in isovelocity knee-extension
Pt immediately after three 3-second isometric
MVCs (+6.1%; p < 0.05), but reported a sig-
nificant decrease in isokinetic knee-extension Pt
immediately after three 5-second isometric
MVCs (3%; p < 0.05). In contrast, Behm et al.[34]
measured isometric MVC peak force after one,
two and three sets of 10-second isometric MVCs,
and the only effect reported was an 8–9%
140
120
Change in twitch torque (%)
100
80
60
40
20
0
−20 Fatigue
protocol
Recovery period
−40
0 2
Time (min)
Fig. 10. The time-course of knee extensor twitch torque during a
fatigue protocol and throughout a subsequent 5-minute recovery
period. The fatigue protocol consisted of 16 5-second MVCs sepa-
rated by 3 seconds of recovery. A twitch contraction was recorded
pre-fatigue protocol, between each MVC, 5 seconds post-fatigue
protocol, and then every 30 seconds throughout the recovery period.
Twitch torque is given as percentages of pre-fatigue values.[14]
ª 2009 Adis Data Information BV. All rights reserved.
Tillin & Bishop
decrease in peak force 10–15 minutes after three
sets of MVCs. As discussed in section 3, PAP is not
expected to enhance isometric peak force (which
represents maximal force), so Behm et al.[34] may
have observed potentiation had they measured
voluntary RFD or dynamic performance. Ad-
ditionally, the smallest CC volume used by Behm
et al.[34] (10-second isometric MVC) is arguably
larger than the smallest CC volume used by
French et al.[50] (three 3-second isometric MVCs
separated by 3 minutes), and may therefore have
induced a greater degree of fatigue. Furthermore,
due to the various other measurements taken
by Behm et al.[34] during the recovery period
(including high-frequency tetanic contractions,
twitches, 30% isometric MVC and ITT), fatigue
may have continued to accumulate, thus reducing
any opportunity to realize PAP.
The results of the four aforementioned stu-
dies[6,14,34,50] demonstrate the influence of CC
volume on the PAP-fatigue relationship. They
also present the possibility that PAP develops
quicker than fatigue and may therefore be uti-
lized immediately after a relatively low CC vo-
lume (window 1 in figure 8). In contrast, as the
CC volume increases so does fatigue and its
dominance in the PAP-fatigue relationship, and
therefore a recovery period may be required be-
fore PAP is realized (window 2 in figure 8). The
specific recovery period required for different
CC volumes is yet to be determined and it is dif-
ficult to compare the results of individual studies
because methodologies have not been standar-
dized. If future research intends to infer the ideal
warm-up and/or training protocol for optimizing
PAP, CC volume and recovery between the
CC and subsequent activity should be assessed
together.
4.3 Conditioning Contraction Type
7 Although, to varying degrees, any type of
contraction is likely to activate the mechanisms of
PAP,[4] the degree of potentiation achieved is
likely to be related to contraction type. Conse-
quently, the use of different types of CC has
probably contributed to the inconsistent results
that have already been discussed. As past research
Sports Med 2009; 39 (2)
Post-Activation Potentiation, Theory and Application
has typically used either isometric or dynamic CC,
this article will only discuss the differences be-
tween these two types of contractions.
Several studies have investigated the effects of
isometric MVCs on subsequent explosive activity,
and whilst two reported an increase,[10,50] others
reported no change in performance.[11,34,57] Past
studies have also used dynamic maximal/near
maximal voluntary contractions to induce PAP,
and again, some recorded potentiation of a
subsequent explosive activity[15,45-48,54,58] and
others did not.[20,49,52,53,55] These conflicting re-
sults (see table I for results) present no clear re-
lationship between contraction type (isometric vs
dynamic) and PAP-response, and only one study
(to our knowledge) has directly compared iso-
metric and dynamic CC with respect to their
effects on performance of a subsequent explosive
activity.[56] This study reported that while a
significant increase in CMJ height (2.9%; p < 0.01)
and peak power (8.7%; p < 0.001) was observed
3 minutes after three 3-second isometric MVC
back-squats, no change in CMJ height (p > 0.05)
but a significant increase in CMJ peak power
(8.0%; p < 0.001) was measured 3 minutes after a
3RM dynamic back-squat set. The authors con-
cluded that their isometric condition induced a
greater PAP-response than their dynamic condi-
tion. The two conditions, however, were not
matched with respect to volume or frequency, and
as a result, it is difficult to make a direct com-
parison between their effects.
Theoretically, different types of contraction
would have different effects on neuromuscular
fatigue.[60,61] Babault et al.[60] assessed neuro-
muscular fatigue during a dynamic contraction
fatiguing protocol and an isometric contraction
fatiguing protocol, where the two protocols were
matched in terms of Pt decrement. The authors
reported that early fatigue during the dynamic
protocol was preferentially peripheral in origin
(peripheral fatigue defined as a decrease in force
generating capacity due to action potential fail-
ure, excitation-contraction coupling failure, or
impairment of cross-bridge cycling in the pre-
sence of unchanged or increased neural drive[61]),
while central fatigue (defined as a reduction in
neural drive to muscle[61]) developed towards the
ª 2009 Adis Data Information BV. All rights reserved.
159
end of the dynamic fatiguing protocol. The iso-
metric protocol, however, produced the opposite
profile, whereby fatigue was firstly central and
then peripheral in origin.
Babault et al.[60] proposed that the difference
in fatigue development between isometric and
concentric contractions might be associated with
muscle metabolite accumulation, which is sug-
gested to activate and/or sensitize groups of small
diameter (III and IV) afferent neural fibres.[60,62,63]
This would in turn cause central fatigue by in-
hibiting a-motoneuron activation, and/or redu-
cing the supraspinal descending drive,[60,63] and/or
decreasing motoneuron firing rate.[64] The inter-
mittent nature of dynamic contractions may fa-
vour blood flow, subsequently aiding the removal
of metabolic by-products. Accordingly, metabo-
lite accumulation would be greater during iso-
metric contractions, resulting in greater central
fatigue. Conversely, lactate accumulation has been
reported to alleviate peripheral fatigue.[65] This
might account for the slower development of per-
ipheral fatigue during isometric contractions
when compared with dynamic contractions.[60]
If isometric and dynamic contractions can in-
duce different fatigue responses, then it is fair to
assume that they might also have different effects
on the mechanisms of PAP. For example, the ec-
centric motion of dynamic contractions (but not
isometric contractions) increases muscle spindle
firing, activating group Ia neural fibres.[63] In
turn, this might enhance the afferent neural volley
at the spinal cord. Consequently, decreased trans-
mission failure from Ia neural fibres to adjacent
a-motor units, resulting in increased higher order
motor unit activation during subsequent activity,
might be greater after dynamic contractions. On
the other hand, isometric contractions activate a
greater number of motor units than dynamic
contractions.[66] Consequently, more muscle fi-
bres might be involved during an isometric
contraction, and this might result in a greater
percentage of RLC phosphorylation, and greater
changes in muscle architecture.
In summary, preliminary evidence suggests
that isometric CCs may induce greater central
fatigue, but are more likely to activate the per-
ipheral mechanisms of PAP. In contrast, dynamic
Sports Med 2009; 39 (2)
160
CCs may induce greater peripheral fatigue, but are
possibly more likely to activate the central me-
chanisms of PAP (table II). The manner in which
these mechanisms interact has not yet been de-
termined, but it is fair to assume that isometric and
dynamic contractions will have different effects on
subsequent explosive activities. The differences
between isometric and dynamic contractions will
also influence the volume and recovery period re-
quired to potentiate subsequent explosive activity.
Future research should investigate the effects of
contraction type on the mechanisms of PAP and
fatigue, whilst standardizing CC volume and re-
covery period. It is also not known whether a CC
of any type is more beneficial than conventional
warm-up methods,[18] and although one study
suggested that it is,[46] their results were specific to
the individuals and protocols assessed. Future re-
search should compare the potentiating effects of
CC to conventional warm-up techniques.
4.4. Subject Characteristics
The subject characteristics that have been
suggested to affect an individual’s PAP-fatigue
response include muscular strength, fibre-type
distribution, training level and power-strength
ratio. These factors are discussed in more detail in
the following sections.
4.4.1 Muscular Strength
There is evidence to suggest that an in-
dividual’s muscular strength might partly de-
termine their PAP response following a CC.
Gourgoulis et al.[15] observed a 4% increase in
CMJ height (p < 0.05) following five sets of back-
squats in those subjects able to squat a load of
>160 kg. Conversely, those subjects unable to
Table II. An illustration of the hypothetical effects of isometric and
dynamic conditioning contractions on the central and peripheral
mechanisms of post-activation potentiation (PAP) and fatigue
Type of The mechanisms of The mechanisms of
conditioning PAP predominantly fatigue predominantly
contraction induced induced
Isometric Peripheral Central
Dynamic Central Peripheral
ª 2009 Adis Data Information BV. All rights reserved.
Tillin & Bishop
squat loads of >160 kg, only recorded a 0.4% in-
crease in CMJ height (p > 0.05). Similarly, Kilduff
et al.[54] reported a correlation between muscular
strength (absolute and relative) and CMJ peak
power potentiation 12 minutes after a 3RM
back-squat set (r = 0.63; p < 0.01). A possible
explanation for these findings might be asso-
ciated with subject fibre-type distribution. The
positive linear relationship between muscular
strength and percentage of type II muscle fibres
is well documented (r = 0.5–0.93; p < 0.05),[67-69]
and type II muscle fibres display the greatest
increase in RLC phosphorylation following a
CC.[7] Furthermore, subjects with a higher per-
centage of type II muscle fibres would pre-
sumably have a greater number of higher order
motor units in reserve, which could be activated
via decreased transmitter failure, following a CC.
The combined effect of a greater RLC phos-
phorylation and a greater increase in higher-
order motor unit recruitment would theoretically
predispose individuals with a higher percentage
of type II muscle fibres to a greater PAP re-
sponse. Consequently, it could be speculated that
the stronger subjects in the two studies discussed
above[15,54] had a higher percentage of fast-twitch
muscle fibres, and thus achieved a greater PAP
response.
4.4.2 Fibre-Type Distribution
Hamada et al.[14] provided evidence to support
a relationship between fibre-type distribution and
PAP. They separated their subjects into two
groups: one with predominantly fast-twitch (type
II) muscle fibres (T-II; n = 4), and a second, with
predominantly slow-twitch (type I) muscle fibres
(T-I; n = 4). They reported a greater Pt response in
the T-II group during a 3-second isometric MVC
(250.0 vs 171.0 N
m; p < 0.01). Furthermore, in
response to a fatigue protocol of 16 5-second
isometric MVCs of the knee extensors, the T-II
group showed significantly greater twitch tension
potentiation during the early stages of the fatigue
protocol (+127% vs +40% increase in Pt after the
third MVC; p < 0.05). However, the T-II group
also had a greater decrease in both twitch Pt and
MVC Pt during the later stages of the fatigue
protocol (p < 0.05). Therefore, although subjects
Sports Med 2009; 39 (2)
Post-Activation Potentiation, Theory and Application
with a greater percentage of type II muscle fibres
elicited a greater PAP response, they also elicited
a greater fatigue response following a high-
volume CC protocol.
There are a number of possible reasons
why Hamada et al.[14] observed a greater fatigue
response in the T-II group. As stated, Hamada
et al.[14] reported a greater Pt production in the T-II
group during the early stages of the fatigue pro-
tocol. Therefore, according to the force-fatigue
relationship,[70] a greater fatigue response in the
T-II group would be expected. Additionally, a
negative correlation has been reported between
initial glycolytic rate and fatigue during inter-
mittent exercise.[71] The specific task employed by
Hamada et al.[14] (16 5-second isometric MVCs,
with 3 seconds of rest between MVCs) would rely
predominantly on a high anaerobic adenosine
triphosphate (ATP) turnover rate, especially in
subjects with a higher percentage of type II mus-
cle fibres.[72,73] Therefore, although subjects with
a higher percentage of type II muscle fibres are
expected to produce a larger MVC Pt, due to a
higher initial anaerobic ATP turnover rate, they
are also likely to show greater Pt decrements, due
to a greater utilization of anaerobic energy stores
and the production of metabolites associated
with fatigue.[74,75]
4.4.3 Training Level
An individual’s training level may also influ-
ence PAP and fatigue responses following a CC.
Chiu et al.[20] separated a sample of 24 subjects
into athletes who were training and participating
in a sport at national and/or international level
(RT; n = 7), and those who participated in re-
creational resistance training (UT; n = 17). Five
sets of one back-squat with 90% 1RM and 5–7
minutes of subsequent recovery induced a 1–3%
increase in CMJ and SJ height in the RT group.
In contrast, the UT group reacted to the same
condition with a 1–4% decrease in CMJ and SJ
height. Chiu et al.[20] suggested that those subjects
training at higher levels of resistance would de-
velop fatigue resistance as an adaptation of their
intensive training regimens, and were more likely
to realize PAP. Chiu et al.,[20] however, did not
ª 2009 Adis Data Information BV. All rights reserved.
161
measure fibre-type distribution, so it is possible
that a greater percentage of fast-twitch muscle
fibres in the RT group also contributed to the
effects observed in this study.
4.4.4 Power-Strength Ratio
There is also evidence to suggest that a sub-
ject’s power-strength ratio will influence their
PAP response to a CC. Schneiker et al.[76] re-
ported a significant negative correlation between
power-strength ratio and potentiation of peak
power during loaded CMJ, executed 2–4 minutes
after one set of 6RM back-squats (r2 = 0.65;
p < 0.05). Furthermore, when the sample of
strength-trained subjects were separated into
those with a power-strength ratio of <19 W/kg
(group 1) and those with a power-strength ratio
of >19 W/kg (group 2), group 1 had a significant
negative correlation between power-strength
ratio and peak power potentiation (r2 = 0.91;
p < 0.05). In contrast, group 2 showed no re-
lationship between power-strength ratio and
peak power potentiation (p > 0.05). These results
suggest that those subjects less able to effectively
convert their strength into power are more likely
to benefit from PAP than those that can. In ad-
dition, it appears that there may be a power-
strength ratio threshold above which subjects do
not benefit from PAP.
In summary, several subject characteristics
have been suggested to affect an individual’s
PAP-fatigue response, and this may partly ex-
plain the inconsistencies of past research. Evi-
dence suggests that individuals most likely to
benefit from PAP include those with a greater
muscular strength, a larger percentage of type
two fibres (although fatigue may also be greater
in these individuals), a higher level of resistance
training, and a smaller power-strength ratio.
Further research, however, is required to validate
these findings as well as determine the possible
effects of other subject characteristics such as
muscle and/or lever lengths. For coaches con-
sidering the implementation of CC prior to ex-
plosive activities (in training or performance), it
may be pertinent to first assess each athlete’s sus-
ceptibility to PAP during the off-season period.
Sports Med 2009; 39 (2)
162
4.5 Type of Subsequent Activity
An additional explanation for the inconsistent
results of past research is the different types of
subsequent explosive activities used to determine
the acute effects of PAP. The types of subsequent
explosive activities employed by previous studies
have included isometric MVCs,[10,34,51] isolated
dynamic contractions (e.g. isovelocity knee
extensions),[11,47,50] and compound ballistic
activities (e.g. CMJ and DJ).[10,15,46,49,52-58] It is
possible that a specific CC will not have the same
effect on different explosive activities.
With regard to differences between isometric
and dynamic explosive contractions, previous
studies have reported moderate to strong corre-
lations between isometric and dynamic RFD
(r = 0.65–0.75),[77] and moderate to strong corre-
lations between isometric and dynamic peak
force (r = 0.66–0.77).[77,78] These results indicate a
clear relationship between tests measuring isometric
and dynamic strength and power. There are,
however, a number of differences in the neural
and mechanical processes involved in isometric
and dynamic activities. For example, the motor
unit recruitment and rate coding for an isometric
contraction will probably be regulated by the size
principle,[79] whereby motor units are recruited in
a hierarchical order of small, followed by higher
order units. On the other hand, dynamic con-
tractions might display a specific pattern of mo-
tor unit recruitment relevant to joint angle and
position through the range of motion.[80] Ad-
ditionally, the eccentric movement involved in
dynamic contractions, but not isometric con-
tractions, would result in a greater afferent
(group Ia neural fibres) input from muscle
spindles.[61,81] As a result, the a-motoneuron ac-
tivation responses for isometric and dynamic
contractions would be different.[82] Furthermore,
utilization of elastic strain energy (stretch-
shortening cycle), stored in the muscles during an
eccentric contraction, provides a significant con-
tribution to overall performance of dynamic
movements.[83-85] The stretch-shortening cycle,
however, is not utilized during an isometric con-
traction and, consequently, isometric contrac-
tions may not reflect the muscles capabilities for
ª 2009 Adis Data Information BV. All rights reserved.
Tillin & Bishop
dynamic situations.[82] Finally, PAP is greatest
whilst the muscle is shortening[86] and extends to
higher stimulation frequencies in concentric
when compared with isometric contractions.[22]
This suggests that PAP may have a performance-
enhancing effect beyond what would be expected
based on isometric contractions.
It is also likely that whilst a specific CC might
enhance performance of a particular dynamic
activity, it might decrease or have no effect on the
performance of a different dynamic activity.
French et al.[50] analysed isovelocity knee exten-
sion, CMJ, DJ and 5-second cycle sprint perfor-
mance before and immediately after three
3-second MVC knee extensions. They reported
significant improvements in DJ height, DJ RFD
and knee extension Pt (+5.0%, +9.5% and +6.1%,
respectively; p < 0.05) after the MVCs, but found
no significant effect in any of the other activities
(p > 0.05). French et al.[50] used time-motion
analysis to explain their results. They reported
that the DJ and knee extension MVC had a
muscle activation period of £0.25 seconds. In
contrast, the CMJ and 5-second cycle sprint had
a muscle activation period of ‡0.25 seconds. Ex-
plosive muscle actions have previously been
defined as those that have an activation period
of £0.25 seconds.[77] French et al.[50] therefore
concluded that PAP was only effective in tasks
defined as explosive muscle actions. The conclu-
sions of French et al.,[50] however, should be
interpreted with caution. Some studies have re-
corded a potentiation effect in CMJ performance,
as well as other activities that otherwise might not
fall under the above definition of explosive mus-
cle action.[10,15,46,51,54,56,58] In addition, French
et al.[50] only measured exercise performance im-
mediately after the CC, and a rest interval may
have been needed for a potentiation effect to be
realised. Finally, the CC exercise was an isola-
tion exercise targeting the knee extensors alone.
The DJ may load the knee extensors to a greater
extent than the CMJ and 5-second cycle sprint,
which would explain the increase in DJ
height/RFD. The CMJ and 5-second cycle sprint,
however, may rely on the contribution of various
other large muscle groups, which due to the
kinematics of the CC, had not been potentiated.
Sports Med 2009; 39 (2)
Post-Activation Potentiation, Theory and Application
These results therefore highlight the importance
of closely matching the kinematics of the CC to
those of the subsequent explosive activity. By
doing so, an individual is more likely to activate
the higher order motor units, phosphorylate the
RLC and change the architecture of those muscle
fibres specifically associated with the subsequent
activity.
The aim of recent research has been to estab-
lish the application of PAP to specific explosive
sports activities. Explosive sports activities are
dynamic in nature so, for the reasons discussed
above, isometric responses to a CC should not be
used to infer effects of the same CC on sub-
sequent sports activities. If researchers are in-
vestigating the application of PAP to a training
scenario, the reported effects of a CC on sub-
sequent ballistic activities (e.g. CMJ and DJ) may
be useful, as ballistic exercises are used in power-
training programmes. On the other hand, whilst
PAP may sometimes be effective in enhancing
performance of a ballistic exercise, it may not
have the same ergogenic effect on performance of
a specific explosive sports activity (e.g. sprinting,
long jump). If PAP is to be utilized in competi-
tion, research must first determine its effects
beyond those reported for ballistic training ex-
ercises. Two recent studies have shown that PAP
can enhance performance of a specific explosive
sports activity, reporting a decrease in sprint time
(-3% over 10 m,[48] -2% over 30 m,[48] and -3%
over 40 m;[45] p < 0.05) 4–5 minutes after the
execution of near maximal (>80% 1RM) back-
squats. Nevertheless, further research is required
to establish the application of PAP to many dif-
ferent explosive sports activities. Furthermore,
even if PAP is consistently shown to enhance
performance of different explosive sports activ-
ities, several practical implications would need to
be addressed to effectively apply PAP to a com-
petitive scenario (such as the need for possible
equipment in the warm-up area and the require-
ment to compete within the optimal recovery
period following activation). As a result of these
impracticalities, the application of PAP to
performance has been challenged,[18] but with
reported increases in performance by >3%,
further investigation is warranted.
ª 2009 Adis Data Information BV. All rights reserved.
163
5. Conclusion
It may be possible to effectively utilize PAP to
enhance mechanical power and therefore perfor-
mance and/or the training stimulus of an explo-
sive sports activity. Evidence over the practical
application of PAP to explosive activities is,
however, inconclusive. The inconsistent results of
past research appear to be due to the complex
interaction of several factors that determine the
degree to which the different mechanisms of PAP
and fatigue are affected. Greater CC volumes and
intensities are expected to induce greater levels of
both PAP and fatigue. However, the rates at
which PAP and fatigue develop and dissipate
may differ, resulting in two windows of oppor-
tunity to potentiate performance; immediately
after a low-volume CC, or after a specific re-
covery period following a high-volume CC. The
type of CC may also have different effects on the
mechanisms of PAP and fatigue. For example,
isometric MVCs may induce central fatigue, but
peripheral PAP, whilst dynamic MVCs may in-
duce the opposite response. The interaction of
these different mechanisms would, in turn, de-
termine the optimal CC volume and recovery
time required to potentiate (if at all) subsequent
performance. Regardless of the above factors, an
individual training at a higher level, with a greater
muscular strength, a greater fast-twitch fibre
distribution and a lower power-strength ratio
may be more likely to benefit from PAP than an
individual without these characteristics. When
interpreting results, consideration should also be
given to the specific application of PAP in sport.
If the intention is to utilize PAP in competition,
only the results of studies reporting the effects of
a CC on performance of a specific explosive
sports activity should be considered. Although
standardization of these various factors provides
future research with an extremely arduous task,
the results of studies showing 2–10% increases in
performance suggests further investigation of
PAP may be worthwhile. It may be pertinent,
however, for research to first establish how the
mechanisms of PAP and fatigue interact under
different conditions before applying PAP to
sport.
Sports Med 2009; 39 (2)
164
Acknowledgements
No sources of funding were used in the preparation of this
review and the authors have no conflicts of interest that are
directly relevant to the contents of the review.
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Correspondence: Mr Neale A. Tillin, School of Sport and
Exercise Science, Loughborough University, Ashby Road,
Loughborough, Leicestershire, LE11 3TU, UK.
E-mail: N.A.Tillin@lboro.ac.uk
Sports Med 2009; 39 (2)